Chromosome Data on Australasian Bryaceae
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J. Hattori Bot. Lab. No. 80: 251 - 270 (Oct. 1996) CHROMOSOME DATA ON AUSTRALASIAN BRYACEAE H.P. RAMSAY1 and J. R. SPENCE2 ABSTRACT. Chromosome numbers in Australasian taxa of the family Bryaceae are now available for eight species of the genus Bryum, for five species of the genus Rosulabryum and for a single species in each of the genera Orthodontium, Brachymenium, Leptobryum and Pohlia from Australia. Data are also given for two species of Bryum from New Zealand and for two species of Rhodobryum from Papua New Guinea. Reported here for the first time in the Southern Hemisphere are chromosome numbers for seven species of Bryum (B. argenteum Hedw. n = lO, B. australe Hampe n=ll (lO + m), B. caespiticium Hedw. n =30, B. clavatum (Schimp.) C. Muell. n = ll (lO + m), B. creberrimum Tay I. n = 30 from Australia; B. blandum J. D. Hook. & Wils. n = 11 and B. sauteri B.S.G. n = 10 from New Zealand). First reports of chromosome numbers are also given for two species of Rosulabryum (R. subtomentosum (Hampe) Spence n = lO, 11 (lO+ m) R. sp. nov. (A) Spence n = 11 (lO + m)) and Pohlia a.ff. nutans (Hedw.) Lindb. n = 22 from Australia while n = 10 is recorded for both Rhodobryum aubertii (Schwaegr.) Ther. and R. giganteum (Schwaegr.) Par. from Papua New Guinea. Additional cytotypes to those previously published were found in several new populations of Brachymenium preissianum (Hampe) Jaeg. [n = 10], for a single population of Bryum pachytheca C. Muell. [n = 20] and for two populations of Orthodontium Jineare Schwaegr. [n = 20] . Additional counts, based on studies of new populations that confirm previous reports for Australian taxa, are given for Bryum dichotomum Hedw. n = 10, Rosulabryum billardierei (Schwaegr.) Spence (Bryum bi/Jardierei Schwaegr. var. bi/Jardierei) n = 10, Rosulabryum sub fasciculatum (Hampe) Spence (Bryum subfasciculatum Hampe) n = 10 and Rosu/abryum torqu escens (Bruch ex De Not.) Spence (Bryum torquescens Bruch ex De Not.) n = 20, 21 (20 + m). INTRODUCTION The family Bryaceae, is one of the largest families of mosses, being characterised by the erect, scarcely branched stems bearing capsules having a more or less elongate sterile neck region and leaf cells rhombic to long vermicular with no specialised alar region. Some genera have been monographed on a worldwide basis (Orthodontium Meijer 1952), while others have been studied only in particular geographical regions. Ochi's taxonomic studies on the family Bryaceae have dealt with the subfamily Bryoideae (1970, 1973, 1979, 1992) and Bryum is the only genus in the family that has been studied to any extent in Australasia (Australia/New Zealand) (Ochi 1970, 1973, 1979). Australian genera in the family are being revised by J. R. Spence & H. P. Ramsay or J. Shaw for inclusion in the Bryophyte volumes (mosses) for the Flora of Australia. 1 School of Biological Sciences, Macquarie University, Sydney 2109 N.S.W. Australia. 2 National Park Service, Glen Canyon National Recreation Area, P.O. Box 1507, Page, Arizona U.S.A. 86040. 252 J. Hattori Bot. Lab. No. 80 I 9 9 6 In Australasia the Bryaceae is a large weedy family favouring open to diffusely shaded habitats, often in disturbed or in marginal ecotonal areas such as road cuts and banks, rock faces, streams (on rock), or open soil. Only a few taxa occur in temperate closed or tropical/subtropical closed forests in Australia where they are found in openings, along streams or on forest margins. Eleven of the fourteen genera and about 90 of the 850 listed species of Bryaceae (Shaw 1985) are present in Australasia including several new or poorly known species. Spence ( 1996) has revised the concept of the genus Bryum and raised most of Section Rosulata in Australasia to generic status as Rosulabryum. Genera represented in this region i.e. Australia and New Zealand (Spence & Ramsay in prep.) are Anomobryum, Bryum, Brachymenium, Epipterygium, Leptobryum, Orthodontium, Plagiobryum, Pohlia, Rhodobryum, Rosulabryum and Schizymenium. CHROMOSOME STUDIES The most common chromosome numbers in the family Bryaceae are n = 10, 11 or 12 and multiples thereof e.g. n = 20, 22, 24, 30, 33, 50, 56 (Fritsch 1991). Previous records of chromosome numbers for Australian Bryaceae (Table 1) can be found in Ramsay (1967, 1974, 1977, 1983) but there are no previous records for New Zealand or Papua New Guinea (included here) species. All vouchers for earlier published numbers have been re-checked and some identifications revised (Table 1). METHODS The present studies have been carried out as part of a long term project on cytological studies of Australasian mosses by Ramsay whose techniques appear in earlier publications ( 197 4, 1983). Meiosis was studied in capsules of Bryaceae using the squash method and staining in aceto-carmine. Slides were made permanent using dry ice and euparal. Having permanent preparations has made it possible to refer back to earlier slides, made as long as 20 years ago, for comparison and checking of counts. For cytological studies, members of the family Bryaceae must be collected and kept in good light to avoid etiolation, although apices of elongated shoots are useful in some cases for mitotic investigation. Plants do not survive for long periods in the laboratory hence care must be taken in selecting material with capsules young enough, but near enough to maturity, to undergo meiosis soon after transfer to the laboratory. The appearance of the capsule at the time of meiosis varied in Bryum from green and translucent with no sign of colour at the tip of the operculum in some, to colouring of the tip of the operculum or capsule wall being indicative of the time of meiosis in others. Seta colour (change from pale green to red/brown just below the capsule) sometimes may be useful in the selection of suitable capsules. Mitotic studies on Rhodobryum aubertii from Papua New Guinea were obtained from premeiotic mitoses in young capsules. Voucher specimens will be deposited at the National Herbarium of New South Wales (NSW). H. P. RAMSAY & I . R. SPENCE: Chromosome data on Australasian Bryaceae 253 Table 1. Previous counts for Australian Bryaceae. Taxon Chrom. No. State Reference Brachymenium preissianum n = 22 Tas Ramsay 1967, 1974 n = 30 W.A. Ramsay 1977 Bryum capi/lare (55/63) n = 20 N.S.W. Ramsay 1967, 1974 Bryum dichotomum n = lO N.S.W. Ramsay 1974 Bryum pachytheca n = lO W.A. Ramsay 1977 Bryum sauteri n = 11 (lO + m) N.S.W. Ramsay 1967, 1974 [ 4 7/ 64 published as B. billardierei redetermined by Spence 1993] Rosulabryum billardierei n = 10, 11 (lO + m) W.A. Ramsay 1977 ( 45/63, 46/63, 67 /64) n = lO, 11 (lO+ m) N.S.W. Ramsay 1967, 1974 [published as Bryum billardierei var. bil/ardierei] Rosulabryum subfasciculatum n = lO Qld Ramsay 1974 [9/65 published as B. bi/lardierei, redetermined by Mohamed (1979)] Rosulabryum torquescens n = 20 N.S.W. Ramsay 1967, 1974 [ 68/64 published as B. billardierei redetermined by Mohamed (1979) as B. torquescens] Rosulabryum torquescens n = 21 (20+ m) Tas Ramsay 1967, 1974 [92/64 published as Bryum billardierei redetermined by Mohamed (1979) as B. torquescens] Rosulabryum torquescens n = 20 W.A. Ramsay 1977 [published as B. campylothecium redetermined by R . Wyatt pers. comm. as B. capi/lare subsp. torquescens] Leptobryum pyriforme n = 22 (21 + m) N.S.W. Ramsay 1974 Orthodontium lineare n = 22 N.S.W. Ramsay 1974 RESULTS The term populations as used here refers to collections made in different localities, often in separate Australian States. Thus they are geographically isolated from each other and could not interbreed. Where specimens occur in the same locality but only a short distance (some metres) apart these are called collections for, although even such a short distance probably precludes interbreeding (sperm transfer distance too far) , they may be genetically related from dispersal by spores or vegetative diaspores. Previous records of chromosome numbers for each of the species examined were checked against Fritsch (1991), who has summarised world literature on chromosome numbers, as well as from publications that have appeared since. 254 J. Hattori Bot. Lab. No. 80 I 9 9 6 Table 2. New Chromosome Number reports for Australasian Bryaceae [* re- fer to Appendix 1 for complete details of locality, date and collector; States are given for Australian collections (Qld, N.S.W., S.A., Vic., W.A.), N.Z. for New Zealand and P.N.G. for Papua New Guinea species]. Species Chrom. No. (n = ) Voucher Numbers Country/State Bryum Section Apa/odictyon B. clavatum 11 (lO + m) 32/77, 33/77 N.S.W. B. sauteri 11 (lO+ m) 91 /84 N.S.W., N.Z. Section Alpiniformia B. australe 11 (lO+ m) 3/88 Tas Section Bryum B. argenteum 10 3/77 Vic., N.S.W. Section Doliolidium B. dichotomum 10 22/73, 27/73 N.S.W. B. pachytheca 10 2/74, 10/76 Vic. 22/77 Section Caespiticia B. caespiticium 20 22/77 Vic. 30 8/77, 9/77 Vic. B. creberrimum 30 44/82, 51 /82, 34/84 N.S.W. , S.A. 6/87 A.C.T. Section Blandum B. blandum 11 76/84 N.Z. Brachymenium B. preissianum 10 52/84, 38/84 S.A. Orthodontium 0 . /ineare 20 51 /84 N.S.W. Pohlia P. aff. nutans 22 40/79 N.S.W. Rhodobryum R. aubertii 10, 2n = 20 16/82, 1/83 P.N.G. R. giganteum 10 9/82 P.N.G. Rosulabryum R. bi/lardierei 10 11/77, 41 /84 N.S.W. 17/72 A.C.T., Vic. R. subfasciculatum 10 11 /72, 26/79 51 /83 N.S.W. R. subtomentosum 10, 11 (lO + m) 25/77' 26/77 54/83, 47/84 N.S.W., Vic.